Effects of defoliation frequency and nitrogen fertilization on the production and potential for persistence of Dactylis glomerata sown in multispecies swards

Management decisions should facilitate the dominance of C₃ perennial grasses over annuals. This study examined the effects of defoliation frequencies and nitrogen fertilization on the productivity and potential for persistence of Dactylis glomerata L. (DG cocksfoot, perennial) in multispecies swards. Treatments were randomly applied to 24 mini‐swards of DG + Bromus willdenowii Kunth (BW prairie grass, annual/biennial) in a factorial design of four defoliation frequencies, based on number of leaves per tiller, by two nitrogen winter fertilization levels (N^? or N⁺). Regardless of fertilization, very frequent and repeated defoliations were related to decreases of about 43% of aboveground biomass and frequent defoliations with decreases of about 44% of vegetative tillers associated with horizontal space occupation and potential for persistence. Nevertheless, differences in DG aerial productivity or reserves were not detected between frequent and optimal defoliation frequencies. Combined effects of N⁺ and optimal frequency were related to root biomass increment of about 200%, compared with frequent defoliation, associated with competitiveness and survival of DG. Optimal defoliation frequency would have ecological but not production advantages, compared with frequent defoliations. The results are discussed in terms of more objective decision‐making in the management of multispecies swards.     

The influence of vernalization on response to nitrogen of perennial ryegrass

Vernalized and unvernalized perennial ryegrass (Lolium perenne cv. Gremie) tillers planted in microplots in a glasshouse in April 1974 and 1975 were exposed to natural daylength thus inducing vernalized plants to produce reproductive, and unvernalized plants vegetative growth. Nitrogen was applied at 0–30 g m^?2 (to 24 g m^?2 in 1975) to both types of herbage. At the first defoliation in each year yields from vernalized grass were greater than those from unvernalized grass; subsequently differences were minimal or reversed. Vernalization had no significant effect (P > 0.05) on total yields over four defoliations in either year. Apart from the first cut in 1974 nitrogen application increased dry matter yields of both types of herbage. Tiller numbers were usually increased by the lowest level of nitrogen application but at higher levels the effect was variable. There was no evidence of an interaction effect of level of nitrogen application and vernalization on dry matter yields.     

The effect of defoliation practice in Western Australia on tiller development of annual ryegrass (Lolium rigidum) and Italian ryegrass (Lolium multiflorum) and its association with forage quality

The effect of defoliation on the vegetative, early reproductive and inflorescence stages of tiller development, changes in the dry‐matter yield of leaf, stem and inflorescence and the associated changes in forage quality was determined on plants of annual ryegrass (Lolium rigidum Gaud.) and Italian ryegrass (L. multiflorum Lam.). The field study comprised seventy‐two plots of 1 m × 2 m, sown with one annual ryegrass and seven Italian ryegrass cultivars with a range of heading dates from early to late; defoliation commenced 6 weeks after germination. During the vegetative stage of growth, plots were defoliated when the tillers had three fully expanded leaves (three‐leaf stage). During the early reproductive stage of growth, to simulate a cut for silage, plots were defoliated 6–7 weeks after 0·10 of the tillers displayed nodal development. The subsequent regrowth was defoliated every 3 weeks. Assessments of changes in tiller density, yield and quality were made in the growth cycle that followed three contrasting cutting treatments during the winter–spring period (from 10 July). In treatment 1, this growth cycle (following closing‐up before a subsequent conservation cut) commenced on 7 August following two defoliations each taken when the tillers were at the three‐leaf stage. In treatment 2, the growth cycle commenced on 16 October following: for early‐maturing cultivars, two cuts at the three‐leaf stage, a cut for silage and an additional regrowth cut; for medium‐maturing cultivars three cuts at the three‐leaf stage and a cut for silage; and late‐maturing cultivars, five cuts at the three‐leaf stage. In treatment 3, defoliation up to 16 October was as for treatment 2, but the growth cycle studied started on 27 November following two additional regrowth cuts for early‐ and medium‐maturing cultivars and cut for silage for the late‐maturing cultivars. Tiller development for all cultivars was classified into three stages; vegetative, early reproductive and inflorescence. In treatment 1, in vitro dry‐matter digestibility (IVDMD) and crude protein (CP) content were negatively associated with maturation of tillers. IVDMD ranged from 0·85 to 0·60 and CP ranged from 200 to less than 100 g kg^–1 dry matter (DM) during the vegetative and inflorescence stages respectively. This large reduction in forage quality was due to an increase in the proportion of stem, inflorescence and dead material, combined with a reduction in the IVDMD and CP content of the stem. A high level of forage quality was retained for longer with later‐maturing cultivars, and/or when vegetative tillers were initiated from the defoliation of early reproductive tillers (treatments 2 and 3). However, 15 weeks after the closing‐up date in treatment 1, defoliation significantly reduced the density of inflorescences with means (±pooled s.e_m.) of 1560, 1178 and 299 ± 108 tillers m^–2, and DM yield of inflorescence with means of 3·0, 0·6 and 0·1 ± 0·15 t ha^–1 for treatments 1, 2 and 3 respectively. This study supports the recommendation that annual and Italian ryegrass cultivars should be classified according to maturity date based on the onset of inflorescence emergence, and that the judicious defoliation of early reproductive tillers can be used to promote the initiation of new vegetative tillers which in turn will retain forage quality for longer.     

The effects of defoliating grass in winter or spring on herbage yields and ensilage characteristics

Under Irish conditions, the digestibility in May of grass managed for silage production is sometimes lower than expected. In each of two successive years, replicate field plots were established to examine the effects of three defoliation heights (uncut or cut to a stubble height of 10 or 5 cm) applied in winter and/or spring on herbage yields harvested in May and again in July, and on chemical composition and conservation characteristics associated with first‐cut silage. Swards that were not defoliated in December or March had a dry‐matter (DM) yield and in vitro DM digestibility (DMD) in mid‐May of 6597 kg ha^?1 and 736 g kg^?1, respectively, in Year 1, and corresponding values of 7338 kg ha^?1 and 771 g kg^?1 in Year 2. Defoliating swards to 5 cm in December reduced (P < 0·001) May DM yields compared to swards that were not defoliated in both December and March, while herbage DMD in May increased (P < 0·001) when defoliated in December or March. There were no clear effects of defoliation height or its timing on herbage ensilability or resultant conservation efficiency characteristics. The effects of defoliation on July yield were the reverse of those observed for May, while the total yield of the December and March defoliations plus the two silage harvests increased as defoliation height was lowered in Year 2 only. It is concluded that defoliation in winter and/or spring can increase herbage digestibility but will likely reduce DM yields in May.     

Changes in the physiology and feed quality of cocksfoot (Dactylis glomerata L.) during regrowth

Abstract A glasshouse study was undertaken to determine the physiological and morphological changes in cocksfoot (Dactylis glomerata L.) during regrowth after defoliation. Individual plants were arranged in a mini‐sward in a randomized complete block design. Treatments involved harvesting each time one new leaf had expanded (one‐leaf stage), up to the six‐leaf stage, with the plants separated into leaf, stubble (tiller bases) and roots. Stubble and root water‐soluble carbohydrate (WSC), stubble and leaf dry matter (DM), tiller number per plant and leaf quality (crude protein (CP), estimated metabolizable energy (ME) and mineral content) were measured to develop optimal defoliation management of cocksfoot‐based pastures. WSC concentration in stubble and roots was highest at the five‐ and six‐leaf stages. Mean WSC concentration (g kg^?1 DM) was greater in stubble than roots (32·7 ± 5·9 vs. 9·4 ± 1·5 respectively). There was a strong positive linear relationship between plant WSC concentration and leaf DM, root DM and tillers per plant after defoliation (Adj R² = 0·72, 0·88 and 0·95 respectively). Root DM plant^?1 and tiller DM tiller^?1 decreased immediately following defoliation and remained low until the three‐leaf stage, then increased from the four‐leaf stage. Tillers per plant remained stable until the four‐leaf stage, after which they increased (from 9·9 ± 0·5 to 15·7 ± 1·0 tillers plant^?1). Estimated metabolizable energy concentration (MJ kg^?1 DM) was significantly lower at the six‐leaf stage (11·01 ± 0·06) than at any previous leaf regrowth stage, whereas CP concentration (g kg^?1 DM) decreased with regrowth to the six‐leaf stage. Both the levels of ME and CP concentrations were indicative of a high quality forage throughout regrowth (11·37 ± 0·04 and 279 ± 8·0 for ME and CP respectively). Results from this study give a basis for determining appropriate criteria for grazing cocksfoot‐based pastures. The optimal defoliation interval for cocksfoot appears to be between the four‐ and five‐leaf stages of regrowth. Delaying defoliation to the four‐leaf stage allows time for replenishment of WSC reserves, resumption of root growth and an increase in tillering, and is before herbage is lost and quality falls due to onset of leaf senescence.     

Herbage yield, tiller number and root system activity after defoliation of prairie grass (Bromus catharticus Vahl)

The effects of defoliation upon root and shoot systems of prairie grass (Bromus catharticus Vahl) were examined in both field and pot studies. The varieties used were 78–32 (HY), a high-yielding variety; 79–42 (LY), a low-yielding variety; and the commercial variety Grasslands Matua. In the field, the presence of roots in early and late spring was estimated by measuring uptake of [³²P]phosphate by roots; herbage yields and tiller numbers were recorded. In a pot study, root and shoot dry-matter (DM) yields were analysed. In the field, roots were detected to a depth of 1·2 m. After defoliation to a height of 0·1 m, root presence decreased more than 50% at depths of 0·6 m for LY and 1·0 m for Matua in early spring and at several depths for each variety in late spring. After a second defoliation, the apparent growth rate of shoots decreased by 35% in relation to the first regrowth period. In pots, shoot DM and root DM of control plants (undefoliated) had the following allometric relationship of the form: In (shoot DM) = 0·61 + 1·14 ln (root DM) (r²= 0·81). After defoliation, compared with undefoliated controls, the relative growth rate of shoots and total herbage yields were higher, but root and stubble DM were lower in all three varieties. Pooled root DM means were 10·3 and 6·8 g plant^?1 and pooled stubble DM means were 12·7 and 7·6 g plant^?1 for control and defoliated plants respectively. HY produced heavier tillers than LY, pooled means being 0·94 and 0·53 g DM tiller^?1 (field study) and 3·44 and 2·05 g DM tiller^?1 (pot study) for HY and LY respectively. HY had 5–6 green leaves per tiller, whereas LY had 3–4. Developed green leaves were heavier in HY (58 g m^?2) than in LY (48 g m^?2). It is suggested that differences in both leaf parameters may be related to higher herbage yields for HY than LY.     

Effects of an autumn defoliation on overwintering, spring growth and yield of a white clover/grass sward

An established sward of binary mixtures of meadow fescue (Festuca pratensis) and white clover (Trifolium repens) (either AberHerald, Grasslands Huia or Sandra) was subjected to (A) no further defoliation, (B) a defoliation in late September or (C) a defoliation in late October after four harvests had been taken during the grazing season. About a tonne of dry matter (DM) was removed by the autumn defoliations. There were two levels of nitrogen application in spring, either 0 or 90 kg ha^?1. The development of grass and clover morphology and population sizes from early autumn until the first harvest the following year was followed by regular sampling of the above-ground material. Stolons were analysed for total non-structural carbohydrates (TNCs), and the temperature at stolon level was continuously recorded. There were no interactions between autumn defoliation, clover cultivar or nitrogen treatments on any of the parameters studied. White clover growing-point numbers and stolon morphological characteristics were reduced in size during the winter and did not recover during the spring. A defoliation in late September resulted in the greatest reduction, whereas there were no differences between the other two treatments. The grass tiller population increased from early autumn until the last sampling occasion in May, but both autumn defoliations resulted in a smaller increase. Defoliation in late September had the greatest impact. The TNC content of white clover stolons fell from about 350 g kg^?1 to 150 g kg^?1 DM from late autumn until late April. There were small differences between the treatments, but a defoliation in late September resulted in a significantly lower level in late autumn. The temperature amplitude at stolon level was consistently greater in plots defoliated in late September. Total DM harvested in spring was 4367, 2564 and 3536 kg ha^?1, of which 388, 352 and 460 kg ha^?1 was white clover, from treatments A, B and C respectively. It is concluded that an autumn defoliation may affect the overwintering of white clover negatively, but that the effect on the grass may be even more detrimental.     

The effect of time of initial defoliation and height of defoliation on the productivity of perennial ryegrass swards

Over a 3-year period the productivity of perennial ryegrass ( Lolium perenne L., cv. S24) swards as influenced by variations in the time of initial defoliation and close (2·5 cm) and lax (8 cm) defoliation was studied. Treatments with initial defoliations in the early vegetative stage, stem elongation and flowering stages of growth gave higher annual yields than treatments in which the initial defoliation was taken at an intermediate stage of growth when the majority of the stem apices had just been elevated above the height of defoliation. Initial defoliation treatments which removed a high proportion of apical meristems while the yield at the initial defoliation was still low produced the lowest annual yield. Consistent lax defoliation reduced yield by 14·5% compared with continual close defoliation. However, taking an initial lax defoliation followed by subsequent close defoliations produced a slight yield advantage of 4·8% over consistent close defoliation. The results are discussed in relation to other research findings and the practical implications commented upon.     

Grazing management and tussock distribution in elephant grass

Soil occupation capacity via lateral expansion of tussocks in elephant grass (Pennisetum purpureum Schum.) may be associated with basal tillering. As grazing management alters the proportion of basal and aerial tillers in a tiller population, the hypothesis of this work was that grazing management affects tussock size and distribution with implications for plant population stability. The objective of this study was to evaluate the tiller population stability index, the proportion of basal and aerial tillers, tussock size, and the frequency of tussocks and bare ground in rotationally managed elephant grass cv. Napier. Treatments resulted from the combination of two post‐grazing heights (35 and 45 cm) and two pre‐grazing conditions (95% and maximum canopy light interception during regrowth – LI_0·95 and LI_Max) and were allocated to experimental units (850 m² paddocks) according to a 2 × 2 factorial arrangement in a randomized complete block design, with four replications. Measurements were taken from January 2011 to April 2012. The post‐grazing height treatments affected the tiller population stability index, but did not influence the pattern of tussock distribution. On the other hand, the different grazing frequencies (targets of LI pre‐grazing) altered the pattern of tussock distribution and the proportion of bare ground. In general, the tiller population stability index and frequency of tussocks were higher and the frequency of bare ground lower on swards managed with the LI_0·95 target relative to those managed with the LI_Max target, regardless of the post‐grazing height used, indicating a larger soil occupation capacity of plants under the more frequent defoliation regime. Such responses were associated with larger population of basal tillers and highlight the importance of tiller category and perennation pathway in defining patterns of plant growth and tussock distribution.     

Leaf number as a criterion for determining defoliation time for Lolium perenne: 2. Effect of defoliation frequency and height

The object of this study was to determine the importance of frequency and height of defoliation on regrowth potential of Lolium perenne. Defoliation interval was based on stage of the regrowth cycle, as indicated by leaves per tiller.
Simulated swards of Lolium perenne cv Yatsyn were grown as individual plants in a glasshouse kept at a day/night temperature of 25°C/15°C.
Treatments imposed were defoliation at 2, 5 or 12 cm residual height, and low and high water soluble carbohydrate (WSC) level obtained by varying defoliation interval, i.e. defoliating at the 1-leaf or 3-leaf stage of the regrowth cycle. Regrowth after frequent short defoliations was only 65% of the less frequently defoliated plants taken over the full regrowth cycle. This was associated with a lower stubble WSC content (2·15 vs 17·5% in stubble) and a twenty-seven-fold difference in the amount of WSC in the stubble per plant. This difference in total WSC was a combined effect of more and heavier tillers and higher WSC content in stubble of plants defoliated less frequently at the end of the regrowth cycle. The regrowth of plants with WSC levels depleted by frequent defoliation when defoliated at 2 cm was significantly below that of those defoliated at 5 and 12 cm.
The results indicate the desirability of defoliating plants at the 3-leaf stage of the regrowth cycle. This not only allows the full regrowth potential to be expressed in that growth cycle, but also in the next cycle, by allowing the replenishment of WSC reserves and optimizing tiller status. The potential to regrow appears then to be based more on the total amount of WSC than the proportion of WSC in stubble.     

The productivity of oats and berseem clover intercrops. I. Primary growth characteristics and forage quality at four densities of oats

Berseem clover (Trifolium alexandrinum L.) was sown as an intercrop with oats (Avena sativa L.) at 0, 30, 60, 90 and 240 oats plants m^?2 in May in 1999 and 2000 in Alberta, Canada. Forage yield and quality were measured at 10‐d intervals between 35 and 88 d after planting (DAP). Yield is defined as the biomass above 6 cm of the soil surface. The dry‐matter (DM) yield of berseem clover in sole crops increased by 5–10 g m^?2 d^?1 between 35 and 55 DAP and then increased by 21–28 g m^?2 d^?1 between 55 and 75 DAP. The DM yields of oats sown at 240 plants m^?2 increased by 26–28 g m^?2 d^?1 over the whole period from 35 to 75 DAP. Oats were the dominant component in the intercrops, even at low densities of oats. Berseem clover grown with 60 oats plants m^?2 received only 0·24 of the incident light when shading by oats peaked at 65 DAP. DM yields of berseem clover in intercrops with 60 oats plants m^?2 averaged 0·14–0·32 of the yields of berseem clover sole crops. Between 35 and 88 DAP in 2000, the crude protein (CP) concentration of berseem clover sole crops declined linearly from 310 to 180 g kg^?1 DM, and the CP concentration of oats exhibited a quadratic response, declining from 350 g kg^?1 DM at tillering to 110 g kg^?1 DM at the soft dough stage. The mean CP concentration of berseem clover in intercrops with 60 oats plants m^?2 was 25 g kg^?1 less than in berseem clover sole crops, indicating that competition by oats reduced the CP concentration of berseem clover. At later sampling dates, CP and DM yields of intercrops with oats at 60 plants m^?2 equalled those with oats at 240 plants m^?2. The addition of berseem clover to oats in intercrops at 60 oats plants m^?2 reduced the neutral‐detergent fibre concentration by 30 g kg^?1 DM compared with oats alone. Oats were very competitive as a companion crop for berseem clover. Adding berseem clover to oats increased forage quality and may provide for increased intake and digestibility of forage to support higher livestock productivity.     

Effect of defoliation management, based on leaf stage, on perennial ryegrass (Lolium perenne L.), prairie grass (Bromus willdenowii Kunth.) and cocksfoot (Dactylis glomerata L.) under dryland conditions. 1. Regrowth, tillering and water-soluble carbohydrate concentration

A field study was undertaken between April 2003 and May 2004 in southern Tasmania, Australia to quantify and compare changes in herbage productivity and water‐soluble carbohydrate (WSC) concentration of perennial ryegrass (Lolium perenne L.), prairie grass (Bromus willdenowii Kunth.) and cocksfoot (Dactylis glomerata L.) under a defoliation regime based on leaf regrowth stage. Defoliation interval was based on the time taken for two, three or four leaves per tiller to fully expand. Dry‐matter (DM) production and botanical composition were measured at every defoliation event; plant density, DM production per tiller, tiller numbers per plant and WSC concentration were measured bimonthly; and tiller initiation and death rates were monitored every 3 weeks. Species and defoliation interval had a significant effect (P < 0·05) on seasonal DM production. Prairie grass produced significantly more (P < 0·001) DM than cocksfoot and ryegrass (5·7 vs. 4·1 and 4·3 t DM ha^?1 respectively). Plants defoliated at the two‐leaf stage of regrowth produced significantly less DM than plants defoliated at the three‐ and four‐leaf stages, irrespective of species. Defoliation interval had no effect on plant persistence of any species during the first year of establishment, as measured by plant density and tiller number. However, more frequent defoliation was detrimental to the productivity of all species, most likely because of decreased WSC reserves. Results from this study confirmed that to maximize rates of regrowth, the recommended defoliation interval for prairie grass and cocksfoot is the four‐leaf stage, and for perennial ryegrass between the two and three‐leaf stages.     

Seed variation among annual ryegrass cultivars in south-eastern USA and the relationship with seedling vigour and forage production

Abstract Annual ryegrass (Lolium multiflorum Lam.) is grown on more than one million ha in the south‐eastern USA each year. Recommended and actual seeding rates vary substantially within the region. The objective of this study was to evaluate variation in seed weight, germination, seedling vigour and seasonal yield performance among annual ryegrass cultivars. During 1997, 1998 and 1999, seed from fourteen commercial cultivars was weighed and germinated to determine numbers of pure live seed (PLS) m^?2 before yield evaluation at four locations. Seed from ten cultivars was planted at 0·7 and 2·0 cm depth in a greenhouse study to evaluate relative seedling vigour. Cultivar mean single‐seed weight ranged from 2·4 to 4·8 mg in 1997, 1·8 to 4·5 mg in 1998, and 2·6 to 4·6 mg in 1999. Seed germination ranged from 78·8% to 98·0% in 1997, 82·3 to 98·3% in 1998 and 77·8 to 98·3% in 1999. Seed number, PLS m^?2, ranged from 675 to 1289 in 1997, 710 to 1550 in 1998, and 717 to 1179 in 1999. Among the ten cultivars evaluated for seedling vigour, seedling weight differed between planting depths and a significant cultivar by year interaction was observed. Seedling weight was highly correlated with seed weight at each seeding depth. The effect of increasing number of PLS m^?2 on subsequent yield performance, although small, was consistently negative. These results indicate that target plant populations may be obtained more economically by adjusting seeding rates for seed size differences among cultivars and seed lots of annual ryegrass.     

FERTILE TILLER PRODUCTION AND SEED YIELD IN MEADOW FESCUE (FESTUCA PRATENSIS L.)

The effect of defoliation and date of nitrogen application in the spring to seed crops of meadow fescue, S215 and S53, were compared in two harvest years, in drills spaced at 18 in. S215 tillers were generally more robust than S53 tillers, and S215 produced a higher number of fertile shoots as the result of a higher conversion rate of over wintering to fertile tillers. N applied early and absence of defoliation gave the highest ratio of seed heads to over wintering tillers.
Seed- and straw-yields were depressed by April and May defoliations; May cutting had a particularly drastic effect on both varieties. S53 yields in all treatments were generally inferior to those of the hay variety, S215.
Head lengths and spikelet numbers were higher and 1000-seed weight lower in S53 than in S215.     

Harvest management based on leaf stage of a tetraploid vs. a diploid cultivar of annual ryegrass

Leaf stage‐dependent defoliation is linked to the plant's physiological status and may be a more suitable criterion than time‐based intervals for harvesting forage grasses, but no reports of research with annual ryegrass (Lolium multiflorum Lam. var. westerwoldicum) were found. To address this, a 2‐year field study was carried out at Raymond, MS, on a Loring silt loam soil (fine‐silty, mixed, thermic Typic Fragiudalfs). Forage production, morphological characteristics and nutritive value responses to defoliation based on leaf stage (2, 3 and 4 leaves per tiller) and two residual stubble heights (RSH; 5 and 10 cm) of a tetraploid (“Maximus”) vs. a diploid (“Marshall”) cultivar of annual ryegrass were quantified. Forage harvested, in 2011, increased linearly as leaf stage increased from 7.3 to 8.8 Mg/ha, but during 2012 was least (7.0 Mg/ha) at 3‐leaf stage and similar at the other two leaf stages (7.6 Mg/ha). Tiller density was less for Maximus (1,191 tillers/m²) than for Marshall (1,383 tillers/m²). Leaf blade proportion decreased with increasing leaf stage and was greater by 9% for Maximus than for Marshall. Generally, forage nutritive value became less desirable with increasing leaf stage. There was a dichotomy in forage harvested and nutritive value responses, but maximum forage productivity was achieved when annual ryegrass was defoliated at the 4‐leaf stage interval.     

THE EFFECTS OF DEFOLIATION ON EMERGENT ANNUAL GRASS SWARDS UNDER SIMULATED GRAZING

For three weeks after emergence, micro-swards of Lolium rigidum were defoliated to a height and at a frequency which represented a range of set stocking rates of less than, similar to, and more than 7–8 sheep/ha as based on the results of Greenwood and Arnold (6) and referred to as D₁, D₂ and D₃ respectively. The effects of these defoliation regimes on the growth and development of emergent swards were evaluated according to their effect on increments of DM from plant parts, rate of appearance of leaves and tillers, concentration of soluble carbohydrates, and on the uptake and concentration of inorganic nutrients. Defoliation began five days after emergence, when the dry weight of the shoot was 3g/m². After 20 days of treatment, 6±7 g/m² of DM had been removed from D₁, 5±5 g/m² from D₂ and 51 g/m² from D₃; and the dry weight of shoot remaining was 13, 7 and 4±5 g/m² on D₁, D₂ and D₃ respectively. The effect of defoliation on dry weight of plant parts increased in severity in the order leaf 1, stem+sheaths, leaf 2, roots, leaf 3, tiller 1, tiller 2, leaf 4. Average relative growth rate of the sward was reduced from 17% per day on D₁ to 13% per day on D₃. There was no necrosis of root tissue. Emergence of the later leaves and of the tillers was retarded by up to three days by more frequent defoliation. Alcohol-soluble carbohydrates in the root fell from over 6% to about 1±5% on all treatments. Uptake of N. P and K was limited by increasing intensity of defoliation and because concentrations of those elements were not greatly affected, the limitation could be ascribed mainly to the effects of defoliation on plant size. The plausibility of the technique for the experimental simulation of grazing of emergent annual grass swards in Western Australia is discussed. It is concluded that for set-stocking rates of less than about 12 sheep/ha, grazing, while reducing DM production just after emergence, is unlikely to produce severe physiological stresses in the sward.     

Priority for allocation of water-soluble carbohydrate reserves during regrowth of Lolium perenne

A glasshouse study was undertaken to determine the priority within the perennial ryegrass (Lolium perenne L.) plant for leaf and root growth and daughter tiller initiation after defoliation, in relation to various levels of water-soluble carbohydrate (WSC) reserves at defoliation. Individual plants were arranged in mini-swards, and underwent varying defoliation frequencies and ambient temperatures before defoliation, and harvest heights at defoliation, to obtain a gradient of WSC content at H₁, the date when all plants were defoliated. Defoliation interval consisted of defoliating either three times at the one new leaf tiller^–1 stage (1-leaf stage) of regrowth, or once only at the 3-leaf stage, up to H₁, while night temperature in the week prior to H₁ was altered from 15°C to either 8 or 20°C. At H₁, plants were defoliated to a stubble height of either 20 or 50 mm. Plants were subsequently destructively harvested at days 4, 6, 8, 12, 18 and 27. Leaf and root extension and tiller dynamics were also measured. On a regrowth timescale, tiller initiation was most sensitive, root regrowth moderately sensitive, and leaf regrowth relatively insensitive to a decrease in WSC. The time of daughter tiller initiation also coincided with replenishment of stubble WSC levels. In contrast to this sequence of regrowth events following defoliation, the quantitative effects on growth were different, with elongation and survival of roots most affected by reduced WSC levels. A 30-fold difference in stubble WSC at H₁ between high and low WSC plants (1·52 vs. 0·05 mg tiller^–1) produced only a 4-fold increase in leaf dry matter (DM) after 27 d (2·2 vs. 0·6 g plant^?1), while tiller number plant^?1 increased 6-fold (138 vs. 23% increase in tiller number from H₁). Root elongation rate was 59 times higher in the high than in the low WSC plants (1·18 vs. 0·02 mm d^?1). From a pasture management perspective, the study confirms that defoliation, coinciding with the 3-leaf stage of regrowth and around a stubble height of 50 mm, optimizes persistence and productivity of perennial ryegrass. By allowing more rapid replenishment of WSC reserves, this optimal defoliation strategy enables a greater proportion of WSC to be allocated to maintain a more active root system, and promotes tillering, compared with more frequent and close defoliation.     

The productivity of oats and berseem clover intercrops. II. Effects of cutting date and density of oats on annual forage yield

Annual forage yields of intercrops of berseem clover (Trifolium alexandrinum L.) and oats (Avena sativa L.), as affected by timing of initial harvest and sowing rate of oats, were investigated. Berseem clover was intercropped with oats at 0, 30, 60, 90 and 240 plants m^?2 of oats in 1999 and 2000 in Alberta, Canada. Cutting date treatments involved initial harvest at 10‐d intervals between 35 and 88 d after planting (DAP), and one or two subsequent harvests of regrowth. Total intercrop dry‐matter (DM) yield averaged 12·9–13·3 t ha^?1 with proportions of 0·21–0·43 of berseem clover in the forage. Treatments with an initial cut at the silage stage of oats (76 DAP in 1999, 88 DAP in 2000, at about soft‐dough stage of oats) had greater yields of oats, lower yields of berseem clover, and lower proportions of berseem clover in total annual yields than treatments with an earlier initial cut. Total crude protein (CP) yield was greater with an initial cut at 65–66 DAP than with a silage‐stage initial cut. With an initial cut at 35 DAP (before stem elongation of oats) or after 65 DAP (after heading of oats), yield potential of oats was sometimes reduced compared with silage‐stage treatments, but this was balanced by a greater yield of berseem clover. The impact of harvest timing on total yield decreased as the proportion of berseem clover in the intercrops increased. With decreasing density of oats, DM yield of first‐cut intercrops and total DM yield of oats decreased, while regrowth and total DM yields of berseem clover increased. Intercrops with oats at 60 plants m^?2 had equal or greater total DM and CP yields than intercrops with 240 plants m^?2 of oats. For intercrops with oats at 60 plants m^?2, with initial cuts at 65–66, 75–76, or 88 DAP, yields of regrowths were 0·30–0·35, 0·16–0·26 and 0·09 of the total yield respectively. Oats–berseem clover intercrops showed potential to manipulate the pattern of annual forage yield and to provide flexibility of harvest without reducing annual yields.     

The impact of defoliation frequency and nitrogen fertilizer application in spring on summer survival of perennial ryegrass under grazing in subtropical Australia

Abstract This field study investigated the effect of timing of nitrogen (N) fertilizer application in spring on the survival of grazed perennial ryegrass (Lolium perenne cv. Dobson and Yatsyn) over summer in a subtropical environment. There were five N fertilizer treatments: no applied N, 46 kg N ha^?1 on 22 October or 22 November or 22 December, or on 22 October and again on 22 December. Water‐soluble carbohydrate (WSC) concentration of perennial ryegrass plants entering the summer was altered by varying defoliation frequency, with defoliation interval based on the number of leaves per tiller. Frequent defoliation was set at a regrowth level of one leaf per tiller and less frequent defoliation at a regrowth level of three leaves per tiller, over a total of two by three‐leaf per tiller regrowth periods. Application of N fertilizer was found to have no significant effect (P > 0·05) on survival of perennial ryegrass plants over summer. On the other hand, defoliation had a marked effect on perennial ryegrass persistence, with frequent defoliation decreasing ryegrass plant density (51 vs. 88 plants m^?2; P < 0·001) and increasing the density of tropical weed grasses (99 vs. 73 plants m^?2; P < 0·001) by autumn. Frequently defoliated plants had a lower stubble WSC content on a per plant basis than less frequently defoliated plants in spring (103 vs. 201 mg per plant; P < 0·001) and summer (59 vs. 101 mg per plant; P < 0·001). The lower WSC content was associated with a smaller root system in spring (1·50 vs. 2·14 g per plant; P < 0·001) and autumn (1·79 vs. 2·66 g per plant; P < 0·01), and this was reflected in 0·29 more plants being pulled from the soil by livestock between November 1996 and April 1997. Rhizoctonia fungus was associated with roots of pulled plants, but not with roots of seemingly healthy plants, indicating that this fungus may have a role in a weakened root system, which was more prone to sod pulling. Nitrogen applied in October and November resulted in a reduced WSC concentration, although the effect was restricted to 1 month after N application. The present study indicates that survival of perennial ryegrass plants over the summer in a subtropical region is prejudiced by frequent defoliation, which is associated with a lower WSC concentration and a shallower root system. Under grazing, sod pulling is a reflection of this weaker root system and contributes to plant mortality.     

Does the “high sugar” trait of perennial ryegrass cultivars express under temperate climate conditions?

The objective was to evaluate water‐soluble carbohydrate (WSC) and crude protein (CP) concentration of perennial ryegrass (PRG) cultivars with different genetic potential for producing WSC under two contrasting agronomic managements in temperate climate (southern Chile). A 4 × 2 factorial design was randomly allocated to 24 plots (31 m² each, three blocks): four PRG cultivars (diploid standard cultivar, “2nSt”; tetraploid standard cultivar, “4nSt”; diploid high sugar cultivar developed in New Zealand, “2nHSNZ”; and tetraploid high sugar cultivar developed in Europe, “4nHSEU”) and two agronomic managements (“favourable,” defoliations at three leaves per tiller and nitrogen (N) fertilization rate of 83.3 kg N ha^?1 year^?1; “unfavourable,” defoliations at two leaves per tiller and N fertilization rate of 250 kg N ha^?1 year^?1). Herbage samples were collected in early spring, spring, summer and autumn. Concentration of WSC did not differ among cultivars in spring and summer, averaging 194 and 251 g/kg DM, respectively. The cultivar 4nHSEU had the greatest WSC concentration in early spring and autumn (187 and 266 g/kg DM, respectively) and the greatest CP concentration across samplings (average 230 g/kg DM). Favourable management improved WSC concentrations in early spring and summer and decreased CP in spring, summer and autumn. Annual DM yield did not vary with cultivar or management, averaging 8.43 t/ha. Within a 12‐month study at one site in a temperate environment in southern Chile, PRG cultivars have not shown a consistent expression of the “high sugar” trait, where a genetic × environment interaction might be operating.